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Research Article
Review of Mesocallis Matsumura from China (Hemiptera, Aphididae), with one new species
expand article infoJing Chen, Li-Yun Jiang, Ge-Xia Qiao§
‡ Institute of Zoology, Chinese Academy of Sciences, Beijing, China
§ University of Chinese Academy of Sciences, Beijing, China
† Deceased author
Open Access

Abstract

The aphid genus Mesocallis Matsumura in China is reviewed. A total of seven species are recognised using morphological characteristics, including six known species, Mesocallis (Mesocallis) alnicola Ghosh, M. (Paratinocallis) corylicola (Higuchi), M. (M.) pteleae Matsumura, M. (M.) sawashibae (Matsumura), M. (P.) yunnanensis (Zhang) and M. (M.) taoi Quednau, and one new species, M. (M.) platycaryae Qiao, sp. nov. The new species, collected on Platycarya strobilacea (Juglandaceae) in Anhui Province, China, is described and illustrated. A key to Mesocallis species from China is presented.

Keywords

Aphids, Calaphidinae, embryo, new host plant

Introduction

The aphid genus Mesocallis was erected by Matsumura (1919), with Myzocallis sawashibae Matsumura, 1917 as the type species. The genus has distinct morphological characteristics; a narrow body, antennae much shorter than the body, antennal segments IV–VI scarcely imbricated, segment III of the alatae with one row of oblong, secondary rhinaria along all or most of its length, and empodial setae distinctly longer than the claws (Higuchi 1972; Quednau 2003; Qiao et al. 2005). Currently, this genus includes 10 species placed in two subgenera: Mesocallis (Mesocallis) alnicola Ghosh, M. (Paratinocallis) corylicola (Higuchi), M. (M.) carpinicola Lee, M. (M.) fagicola Matsumura, M. (M.) obtusirostris Ghosh, M. (P.) occulta Lee, M. (M.) pteleae Matsumura, M. (M.) sawashibae (Matsumura), M. (P.) yunnanensis (Zhang), and M. taoi Quednau (Blackman and Eastop 2020; Favret 2020). These species all are associated with plants of the family Betulaceae and are mainly distributed in East Asia. Six species are hitherto recorded from China: M. alnicola, M. corylicola, M. pteleae, M. sawashibae, M. taoi, and M. yunnanensis. Recently, some apterous specimens on Platycarya strobilacea (Juglandaceae) were collected in Anhui Province (Dabieshan Mountain), which were identified as a new species in this genus. Herein, the genus Mesocallis from China is reviewed, a key to Chinese species is provided, and the new species is described and illustrated.

Materials and methods

The brief procedure of making aphid slide-mounted specimens follows that of Jiang et al. (2016). The descriptions and drawings provided here were produced from slide-mounted specimens using a Leica DM4000B and drawing tube. The photomicrographs were prepared with a Leica DM2500 using DIC illumination and processed with Automontage and Photoshop software.

Aphid terminology in this paper generally follows that of Quednau (2003) and Qiao et al. (2005). The unit of measurements is millimetres (mm). All specimens, including the holotype and paratypes, are deposited in the National Zoological Museum of China, Institute of Zoology, Chinese Academy of Sciences, Beijing, China (NZMC).

Taxonomy

Mesocallis Matsumura, 1919

Mesocallis Matsumura 1919: 103. Type species: Myzocallis sawashibae Matsumura, 1917; by original designation.

Subgenus Mesocallis Matsumura 1919: 103

Synonym: NeocallisMatsumura 1919: 104.

Synonym: NippochaitophorusTakahashi 1961: 247.

Subgenus Paratinocallis Higuchi 1972: 30. Type species: Paratinocallis corylicola Higuchi,1972; by original designation. Given subgenus status by Quednau (2003: 20).

Mesocallis Matsumura: Higuchi 1972: 22; Raychaudhuri et al. 1980: 291; Ghosh and Quednau 1990: 114; Quednau 2003: 19; Qiao et al. 2005: 194; Lee et al. 2018: 3.

Generic diagnosis

In alatae, eyes with ocular tubercles. Antennae 6-segmented, processus terminalis 0.80–1.20× as long as the base of the segment. Ultimate rostral segment with 2–16 accessory setae. First tarsal segments with five to seven ventral setae and two dorsal setae. Empodial setae flabellate. Siphunculi truncated, without flange, without any setae surrounding at base. Cauda knobbed. Anal plate bilobed. Gonapophyses fused, with eight gonosetae. In apterae, nymphs and embryo, dorsal body setae with spinulose shafts, and round knobs at apex. Abdominal tergites I–IV with one pair of marginal setae in subgenus Mesocallis, or two or three pairs of marginal setae in subgenus Paratinocallis. Compound eye of apterous morph often smaller and with fewer facets than in the alate morph, and inner setae of antennal segment III inconspicuous. Apterae with 5- or 6-segmented antennae, dorsal setae of tibiae similar to other tibial setae in subgenus Mesocallis, or strongly differentiated from other tibial setae in subgenus Paratinocallis; first tarsal segments with five ventral setae, without dorsal setae. In embryo, dorsal body setae capitate at apex; spinal setae of metanotum and tergites I, III, and V short or minute, pleural setae absent. Viviparae alate and apterous in some species.

Distribution

China, Japan, Korea, and India.

Host plants

Alnus, Carpinus, Corylus, and Ostrya (Betulaceae), and Platycarya (Juglandaceae).

Comments

Of the known Myzocallis species, most infest plants of Betulaceae. Two species (obtusirostris and taoi) are primarily associated with Alnus; and three species (corylicola, occulta, and yunnanensis) are associated with Corylus. Myzocallis carpinicola is recorded only on Carpinus. Additionally, M. alnicola infests both Alnus and Corylus, and M. sawashibae occurs on both Carpinus and Corylus (Lee et al. 2018). Only M. pteleae infests plants in several different genera of Betulaceae (Alnus, Betula, Corylus, Carpinus, and Ostrya) (Holman 2009). So, Mesocallis species have distinct host specialization. However, the new species, M. platycaryae Qiao was found on Platycarya (Juglandaceae). All species occur only in East Asia and are endemic to this region.

Mesocallis (Mesocallis) alnicola Ghosh, 1974

Mesocallis alnicola Ghosh 1974: 425.

Mesocallis alnicola Ghosh: Raychaudhuri et al. 1980: 292; Ghosh and Quednau 1990: 116; Quednau 2003: 20; Qiao et al. 2005: 195.

Specimens examined

Two alate viviparous femlaes, China: Gansu (Yuzhong County: Xinglong Mountain, alt. 2300 m), 1 Aug. 1986, no. 8579, on Corylus heterophylla, coll. G.X. Zhang, J.H. Li, and T.S. Zhong (NZMC).

Distribution

China (Gansu), India.

Host plants

Corylus heterophylla in China (first record from this host), Alnus nepalensis in India.

Biology

Yellow in life. Infesting the underside of leaves of host plants.

Mesocallis (Paratinocallis) corylicola (Higuchi, 1972)

Paratinocallis corylicola Higuchi 1972: 30; Qiao et al. 2005: 210.

Mesocallis (Paratinocallis) corylicola (Higuchi): Quednau 2003: 21; Lee et al. 2018: 8.

Specimens examined

Four alate viviparous females and 2 nymphs, China: Heilongjiang (Harbin City), 27 Jul. 1976, no. 6423, on Corylus heterophylla, coll. G.X. Zhang and T.S. Zhong (NZMC); 3 alate viviparous females, Liaoning (Shenyang City), 25 May 1984, no. Y4994, on Corylus heterophylla, coll. L.J. Liu and Y.Q. Wang (NZMC); 2 alate viviparous females, Liaoning (Dandong City), 22 Jun. 1984, no. Y4914, on Corylus heterophylla, coll. G.X. Zhang and L.J. Liu (NZMC); 7 alate viviparous females, Shandong (Taian City), 12 Jun. 1975, no. 5990, on Corylus heterophylla, coll. T.S. Zhong (NZMC); 2 alate viviparous females, Gansu (Yuzhong County: Xinglong Mountain, alt. 2170 m), 30 Jul. 1986, no. 8556, on Corylus heterophylla, coll. G.X. Zhang, J.H. Li, and T.S. Zhong (NZMC).

Distribution

China (Liaoning, Heilongjiang, Shandong, Gansu), Japan, Korea.

Host plants

Corylus sieboldiana and C. heterophylla; however, in the Russian Far East it was collected from Quercus dentata (Holman 2009).

Biology

Beige or pale green in life; scattered on the underside of leaves of host plants.

Mesocallis (Mesocallis) platycaryae Qiao, sp. nov.

Figures 1–12, 13–15, 16–24, Table 1

Etymology

The specific name platycaryae is based on the host plant (Platycarya) of the species.

Description

Apterous viviparous female: body oval (Fig. 13), translucent white in life. For morphometric data, see Table 1.

Table 1.

Morphometric data for apterous viviparous females and nymphs of Mesocallis (Mesocallis) platycaryae sp. nov.

Characters Apterous viviparous female (holotype) Apterous viviparous female (paratype) 3rd nymph (n = 1) 4th nymph (n = 1)
Body length 1.120 1.010 1.020 1.030
Body width 0.470 0.450 0.490 0.490
Antenna 0.532 0.470 0.465 0.483
Antennal segment I 0.047 0.040 0.045 0.042
Antennal segment II 0.037 0.035 0.037 0.042
Antennal segment III 0.183 0.166 0.151 0.144
Antennal segment IV 0.106 0.092 0.087 0.097
Base of antennal segment V 0.084 0.077 0.082 0.089
Processus terminalis 0.074 0.062 0.064 0.069
Ultimate rostral segment 0.062 0.062 0.062 0.062
Hind femur 0.260 0.230 0.225 0.218
Hind tibia 0.396 0.361 0.342 0.302
Second hind tarsal segment 0.084 0.079 0.079 0.077
Siphunculus 0.037 0.037 0.027 0.030
Basal width of siphunculus 0.054 0.050 0.050 0.054
Distal width of siphunculus 0.032 0.035 0.032 0.030
Cauda 0.099 0.099
Basal width of cauda 0.089 0.094
Basal diameter of antennal segment III 0.015 0.012 0.012 0.015
Widest width of hind femur 0.052 0.050 0.050 0.050
Width of hind tibia at mid length 0.020 0.020 0.027 0.025
Longest dorsal cephalic seta 0.084 0.074 0.074 0.057
Longest marginal seta on abdominal tergite I 0.079 0.114 0.059 0.040
Longest seta on abdominal tergite VIII 0.151 0.129 0.092 0.094
Longest seta on antennal segment III 0.005 0.005 0.005 0.005
Longest seta on hind tibia 0.027 0.030 0.040 0.030

Mounted specimens

Body dorsum pale; antennae, legs, cauda, anal plate, and genital plate pale, apex of rostrum brown (Fig. 13). Dorsal body setae thick long and dark brown, with elevated bases, sparsely spinulose shafts on part of length and large round knobs at apices (Figs 4–8, 13, 20–22, 24).

Head. Frons convex (Figs 1, 17). Dorsal setae on head similar to dorsal body setae. Head with one pair of frontal setae (Figs 4, 16, 17), one pair of setae between antennae and two pairs of posterior marginal setae between eyes (Figs 1, 16, 17). Frontal setae 5.67–6.00× as long as basal diameter of antennal segment III. Eyes with relatively few facets (Figs 1, 16). Antennae 5-segmented, 0.29–0.33× as long as body (Figs 2, 18), segments III–V with spinulose transverse imbrication; processus terminalis 0.81–0.88× as long as the base of the segment. Antennal setae very few short and pointed; segments I–V each with 2, 2, 2 or 3, 2, 1 setae, respectively; processus terminalis with four apical setae. Length of setae on segment III 0.33–0.40× as long as basal diameter of the segment. Primary rhinaria not ciliated (Figs 2, 18). Rostrum (Figs 3, 17) reaching back to between fore and mid-coxae; ultimate rostral segment thick wedge-shaped, 1.39–1.56× as long as its basal width, 0.74–0.78× as long as second hind tarsal segment, with two accessory setae

Figures 1–12. 

Mesocallis platycaryae sp. nov. Apterous viviparous female 1 dorsal view of head 2 antennal segments I–V 3 ultimate rostral segment 4 frontal seta 5 marginal seta on abdominal tergite I 6 spinal seta on abdominal tergite III 7 marginal seta on abdominal tergite III 8 spinal seta on abdominal tergite VIII 9 siphunculus 10 cauda 11 anal plate 12 subgenital plate. Scale bars: 0.05 mm.

Thorax (Fig. 13). Pronotum with 1 pair of short, pale brown, anterior spinal setae and 2 pairs of thick, long, dark brown marginal setae; meso- and metanotum each with one pair of spinal and one pair of marginal thick, long, dark brown setae. Mesosternal furca separated (Fig. 5). Femur and trochanter partially fused; hind femur and trochanter 4.66–5.00× as long as greatest width of segment; 1.42–1.48× as long as antennal segment III. Distal half of tibiae and tarsi with spinulose transverse striae (Fig. 19); hind tibia 0.35–0.36× as long as body. Setae on legs fine, pointed; tibial distal setae similar to other tibial setae; length of setae on hind tibiae 1.38–1.50× as long as middle diameter of segment. First tarsal segments each with five ventral setae, and without dorsal setae. Second hind tarsal segment 1.13–1.28× as long as processus terminalis, 0.79–0.86× as long as antennal segment IV. Empodial setae flabellate.

Figures 13–15. 

Mesocallis platycaryae sp. nov. Dorsal view of body 13 apterous viviparous female 14 4th nymph 15 3rd nymph. Scale bars: 0.1 mm.

Abdomen. Abdominal tergites I–VII each with a single pair of spinal and a pair of marginal setae (Figs 5–7, 13, 20–22), marginal setae on tergites I and V distinctly shorter than ones on tergites II–IV (Figs 20, 21), setae on tergite VII very much shorter and pointed, not elevated at base; sometimes spinal setae on tergites III and V slightly shorter than other spinal setae; tergite VIII with one pair of thick long and dark brown dorsal setae (Figs 8, 24). Marginal setae on tergite I are 5.33–9.20× as long as basal diameter of antennal segment III; dorsal setae on tergite VIII 10.17–10.40× as long as basal diameter of antennal segment III. Siphunculi truncated (Figs 9, 22, 23), 0.68–0.75× as long as their basal widths, 0.38× as long as cauda. Cauda knob-shaped, with spinulose short striae (Figs 10, 24); 0.27–0.28× as long as its basal width, with 6–8 long and short pointed setae. Anal plate bilobed, with short spinulose striae (Figs 11, 24). Subgenital plate transversely oval (Fig. 12), with sparse spinulosity in transverse lines; with nine anterior setae, six to eight posterior setae. Gonapophyses fused, with eight short gonosetae.

Figures 16–24. 

Mesocallis platycaryae sp. nov. Apterous viviparous female 16 dorsal view of body, dorsal setae shown 17 dorsal view of head, antennal segments I–II, and ultimate rostral segment 18 antenna, fore tibia and tarsal segment 19 hind tibia and tarsal segments 20 marginal setae on antennal segments I–IV 21 marginal setae and marginal setae on abdominal tergites IV–V 22 siphunculi, spinal and marginal setae on abdominal tergites VI–VIII, cauda and anal plate 23 siphunculus 24 dorsal setae on abdominal tergite VIII, cauda, and anal plate. Scale bars: 0.05 mm.

Third instar nymph. Body oval (Fig. 14), pale brown. Cauda circular at apex, otherwise similar to apterous viviparous female.

Fourth instar nymph. Body oval (Fig. 15), pale brown. Cauda circular at apex, otherwise similar to apterous viviparous female.

Embryo. Dorsal body setae thick, long, and with terminal large round knobs. Head with three pairs of anterior dorsal setae, and two pairs of posterior marginal setae; pronotum with two pairs of spinal setae and one pair of marginal setae, some anterior spinal setae minute; meso- and metanotum each with one pair of spinal and one pair of marginal setae; abdominal tergites I–VII each with one pair of spinal and one pair of marginal setae; among spinal setae of metanotum and tergites I, III, and V are minute, marginal setae on tergites I–III and V–VII; spinal setae on tergites III, V, and VII are displaced.

Specimens examined

Holotype: apterous viviparous female, China (Anhui Province: Yuexi County, Yaoluoping Reserve, Xiaoqiling, alt. 1100 m), 19 Jul. 2007, no. 20714-1-1-1, on Platycarya strobilacea, coll. J.J. Yu (NZMC). Paratypes: 1 apterous viviparous female, 1 third instar nymph, and 1 fourth instar nymph (NZMC), the collection data is the same as in the holotype.

Taxonomic notes

Based on the following morphological characteristics in apterae and nymphs of dorsal body setae with round knobbed apex, 5-segmented antennae, much shorter than the body, hind tibial distal setae similar to other setae on the segment, distal part of tibiae and tarsi spinulose, and abdominal tergites I–VII each with one pair of marginal setae, the new species should clearly be placed in Mesocallis. The species is characterised by the dark-brown dorsal body setae, which are placed on unsclerotized tuberculate bases, and by its colonisation of Platycarya strobilacea (Juglandaceae). Mesocallis platycaryae resembles M. taoi in the number of antennal segments, the ratio of antennae to body length, the length and the number of accessory setae of ultimate rostral segment etc., but apterae differ from those of M. taoi as follows: dorsal body setae dark brown, not arising from sclerites (M. taoi has the dorsal body setae pale but on pigmented sclerites); shafts of dorsal body setae largely smooth, only sparsely spinulose on part of length (vs long dorsal body setae with spinulose shafts); antennae and tarsi pale (vs distal part of antennal segments III–V and tarsi brown). The new species differs from M. carpinicola and M. pteleae in: ultimate rostral segment 0.06 mm long, and with two accessory setae (M. carpinicola and M. pteleae: 0.10–0.14 mm long, with four or more accessory setae), head vertex and antennal segments I–III pale (vs blackish), cauda with 6–8 setae (vs 7–15 setae). In addition, the new species may be distinguished from M. obtusirostris by: antennae 0.46–0.48× as long as body (M. obtusirostris: antennae 0.61–0.75× as long as body), ultimate rostral segment 0.74–0.78× as long as second hind tarsal segment (vs 0.50–0.55×). The difference between the new species and other species of subgenus Mesocallis may be found in the key below.

Distribution

China (Anhui).

Host plant

Platycarya strobilacea (Juglandaceae).

Biology

The species lives scattered on the underside of leaves of host plant.

Mesocallis (Mesocallis) pteleae Matsumura, 1919

Mesocallis pteleae Matsumura 1919: 103. Higuchi 1972: 23; Quednau 2003: 20; Qiao et al. 2005: 196; Lee et al. 2018: 4.

Agrioaphis hashibamii Shinji 1935: 287; Tseng and Tao 1938: 209; Tao 1963: 57.

Myzocallis colyricola Shinji 1941: 1148.

Specimens examined

Two alate viviparous females, China: Hebei (Wulin Mountain), 15 Jul. 1983, no. Y4357, on Corylus mandshurica, coll. S.B. Tian (NZMC); 1 alate viviparous female, Hebei (Wulin Mountain), 13 Sep. 1983, no. Y4354, on Corylus heterophylla, coll. S.B. Tian (NZMC); 1 alate viviparous female, Gansu (Yuzhong County: Xinglong Mountain, alt. 2170m), 30 Jul. 1986, no. 8556, on Corylus heterophylla, coll. G.X. Zhang, J.H. Li, and T.S. Zhong (NZMC); 3 alate viviparous females, Gansu (Tianshui City: Maijishan Mountain, alt. 1700 m), 24 Jul. 1985, no. 8556, on Corylus heterophylla, coll. G.X. Zhang and T.S. Zhong (NZMC).

Distribution

China (Hebei, Sichuan, Gansu), Japan, Korea.

Host plants

Corylus heterophylla, C. sieboldiana var. mandshurica, Alnus cremastogyne in China; but, in Japan, Alnus matsumurae, Corylus heterophylla var. thunbergii (Shinji 1935); C. sieboldiana, C. sieboldiana var. mandshurica (Higuchi 1972), also recorded from Ostrya japonica and Carpinus sp. (Quednau 2003), and Betula spp. (Holman 2009).

Biology

Pale green in life; scattered on the underside of leaves of host plants.

Mesocallis (Mesocallis) sawashibae (Matsumura, 1917)

Myzocallis sawashibae Matsumura 1917: 374; Matsumura 1919: 103.

Mesocallis sawashibae (Matsumura): Higuchi 1972: 24; Quednau 2003: 20; Qiao et al. 2005: 198; Lee et al. 2018: 8.

Neocallis carpinicola Matsumura 1919: 105.

Nippochaitophorus moriokaensis Takahashi 1961: 247.

Specimens examined

Nine alate viviparous females, China: Hebei (Changli County), 30 May 1984, no. 5518, on Corylus heterophylla, coll. S.B. Tian (NZMC); 1 alate viviparous female, Hebei (Wulin Mountain), 15 Sep. 1983, no. Y4357, on Corylus mandshurica, coll. S.B.Tian (NZMC).

Distribution

China (Hebei), Japan, Korea.

Host plants

Corylus heterophylla and C. mandshurica in China; in Japan, Corpinus cordata (Higuchi 1972), Corpinus erosa, C. coreana (Quednau 2003).

Biology

White in life; scattered on the underside of leaves of host plants.

Mesocallis (Mesocallis) taoi Quednau, 2003

Mesocallis taoi Quednau 2003: 20, 53.

Distribution

China (Sichuan).

Host plants

Alnus cremastogyne.

Mesocallis (Paratinocallis) yunnanensis (Zhang, 1985)

Paratinocallis yunnanensis Zhang 1985: 220; Qiao et al. 2005: 211.

Mesocallis (Paratinocallis) yunnanensis (Zhang): Quednau 2003: 21.

Specimens examined

Three alate viviparous females, China: Yunnan (Lijiang City: Yulongxueshan Mountain), 30 May 1984, no. 7192, on Corylus heterophylla, coll. T.S. Zhong (NZMC).

Distribution

China (Yunnan).

Host plants

Corylus heterophylla.

Biology

Beige in life; infesting the underside of leaves of host plants.

Key to the species of Mesocallis in China

1 Abdominal tergites I–IV each with one pair of marginal setae 2
Abdominal tergites I–IV each with two or three pairs of marginal setae 6
2 Dorsal body setae dark brown; shafts of setae mainly smooth, only sparsely spinulose for part of length; on Platycarya strobilacea (Juglandaceae) M. platycaryae sp. nov.
Dorsal body setae unpigmented; with spinulose shafts; on plants of Betulaceae 3
3 In the alatae anterior part of head black; antennal segment III black; ultimate rostral segment 0.7–1.4× as long as hind second tarsal segment 4
In the alatae anterior part of head pale; antennal segment III black in whole, or dorsal half, or only apex; ultimate rostral segment 0.6–0.9× as long as hind second tarsal segment 5
4 In alatae: processus terminalis 0.6–0.8× as long as the base of the segment; ultimate rostral segment 0.7–0.9× as long as hind second tarsal segment; first tarsal segments with five ventral setae; in apterae and nymph: marginal setae of abdominal tergites V and VII minute or very short M. taoi 1
In alatae: processus terminalis 0.9–1.2× as long as the base of the segment; ultimate rostral segment 1.2–1.4× as long as hind second tarsal segment; first tarsal segments with six ventral setae; in alatoid nymph: marginal setae of abdominal tergites V and VII slightly shortened than those on other tergites M. pteleae
5 Some abdominal tergites with duplicated spinal setae and/or with an intercalary seta developed amidst spinal setae; processus terminalis 0.6–0.7 × as long as the base of the segment; in alatae antennal segment III black except for its very base, antennal segment IV sometimes with secondary rhinaria M. alnicola
Abdominal tergites each with one pair of spinal setae; processus terminalis 1.0–1.2× as long as the base of the segment; in alatae antennal segment III pale with apex, antennal segment IV without secondary rhinaria M. sawashibae
6 In alatae: processus terminalis 0.9–1.0× as long as the base of the segment; antennal segment IV with secondary rhinaria; ultimate rostral segment 0.9–1.0× as long as hind second tarsal segment M. corylicola
In alatae: processus terminalis 0.6–0.7× as long as the base of the segment; antennal segment IV without secondary rhinaria; ultimate rostral segment 1.5–1.7× hind second tarsal segment M. yunnanensis

Acknowledgements

We are very grateful to all collectors for their efforts in capturing aphid specimens, and we thank F.D. Yang for helping to make the slide mounts. The work was supported by the National Natural Sciences Foundation of China (grant no. 31620103916), National Science & Technology Fundamental Resources Investigation Program of China (grant no. 2019FY101800), the Key Collaborative Research Program of the Alliance of International Science Organizations (grant no. ANSO-CR-KP-2020-04), and the Youth Innovation Promotion Association of Chinese Academy of Sciences (grant no. 2020087).

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1 The information on M. taoi is based on the original description by Quednau (2003).